Wood anatomy of Mollinedia glabra (Spreng.) Perkins

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Anais da Academia Brasileira de Ciências (2010) 82(4): 915-924(Annals of the Brazilian Academy of Sciences)ISSN 0001-3765www.scielo.br/aabc

Wood anatomy of Mollinedia glabra (Spreng.) Perkins (Monimiaceae)in two Restinga Vegetation Formations at Rio das Ostras, RJ, Brazil

FERNANDA DA S. NOVAES1, CÁTIA H. CALLADO2,MARIA VERÔNICA L. PEREIRA-MOURA1 and HELENA R.P. LIMA1

1Universidade Federal Rural do Rio de Janeiro, Instituto de Biologia, Departamento de BotânicaRod. BR 465, km 7, Caixa Postal 74582, 23890-000 Seropédica, RJ, Brasil

2Universidade do Estado do Rio de Janeiro, Instituto de Biologia Roberto Alcântara GomesDepartamento de Biologia Vegetal, Laboratório de Anatomia Vegetal

Rua São Francisco Xavier, 524-PHLC, 20550-013 Rio de Janeiro, RJ, Brasil

Manuscript received on November 3, 2009; accepted for publication on July 1, 2010

ABSTRACT

This paper aimed to characterize the anatomical structure of the wood of specimens of Mollinedia glabra (Spreng.)

Perkins growing in two contiguous formations of restinga vegetation at Praia Virgem, in the municipality of Rio das

Ostras, RJ. Both the Open Palmae (OPS) and the Sandy Strip Closed Shrub (SSCS) formations are found in coastal

regions that receive between 1,100 and 1,300 mm of rainfall per year. Sapwood samples were collected in both

formations. Typical anatomical features for this species include: solitary vessels, radial multiples or clusters elements,

that are circular to angular in outline, 5-15 barred scalariform perforation plates, wood parenchyma scanty, septate

fiber-tracheids, and wide multiseriate rays with prismatic crystals. Statistical analyses indicated a significant increase

in the frequency of vessel elements and an increase in fiber-tracheid diameters in OPS individuals. These characteristics

are considered structural adaptations to increased water needs caused by a greater exposure to sunlight. Continuous

pruning may be responsible for the tyloses observed in OPS plants. The greater lengths and higher frequencies of the

rays in SSCS trees may be due to the greater diameters of their branches. Our results suggest that M. glabra develops

structural adaptations to the restinga micro-environmental variations during its development.

Key words: Mollinedia glabra, Monimiaceae, restinga, wood anatomy.

INTRODUCTION

Monimiaceae (sensu stricto) is a Pantropical familycomprising 25 to 30 genera and about 200 species. Fivegenera are cited for Brazil: Grazielanthus Peixoto &Per.-Moura, Hennecartia Poisson; Macropeplus Per-kins; Macrotorus Perkins, and Mollinedia Ruiz & Pav(Peixoto and Pereira-Moura 2008).

The genus Mollinedia has the largest number ofdescribed species and is typified by highly branchedshrubs or small trees with erect trunks (Peixoto 1979)that normally grow near rivers and streams. The wood

Correspondence to: Helena Regina Pinto LimaE-mail: [email protected]

of this taxon is hard, whitish, and easy to work, al-though it is little harvested commercially (Peixoto et al.2001). These trees have medicinal uses (Leitão et al.1999) and some species are used in urban landscapingand to produces barrels and sieves in southern Brazil,they also have an important role in reforestation andin the maintenance of the local fauna (Peixoto et al.2001). Mollinedia glabra (Spreng.) Perkins is consid-ered a threatened species in Brazil (Ibama 2008), withsmall populations that occur only in coastal lowlands.The species occurs in two distinct vegetation formationsin the restinga region of Praia Virgem, municipality ofRio das Ostras, Rio de Janeiro: Open Palmae Shrub(OPS) and Sandy Strip Closed Shrub (SSCS).

An Acad Bras Cienc (2010) 82 (4)

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916 FERNANDA DA S. NOVAES et al.

Most studies concerning this genus in Brazil have

focused on their taxonomic (Peixoto et al. 2001, 2002)

or phytochemical (Leitão et al. 1999) nature, with only

infrequent works concentrating on the anatomical or

ecological aspects of this taxon (Barros et al. 1997,

Callado et al. 1997). Anatomical studies of wood are

considered to be very informative (Metcalfe 1989),

mainly when correlated with evolutive and phylogen-

etic questions, as environmental factors are considered

active in defining the structure of the secondary xylem

(Aguilar-Rodríguez and Barajas-Morales 2005, Carl-

quist 2000). Although the first anatomical studies con-

sidering ecological aspects of plant populations were

first performed in the 1970’s (Baas 1973, Carlquist

1975), this type of investigation is still uncommon in

the tropics – largely due to the great variety of species

and environments found there. In general, researchers

that use the anatomical characters of wood in studies

of intra-specific variation in ecologically distinct envi-

ronments have described modifications in the diameter,

length, and frequency of the vessel elements, the thick-

ness of the walls and the lengths of the fibers, and the

height and width of the rays (Baas et al. 1983, Callado

et al. 1997, Wheeler and Baas 1991, Marcati et al. 2001,

León 2005, Ribeiro and Barros 2006).

The present work described and compared the

wood anatomy of individuals of M. glabra growing in

two contiguous vegetation formations under distinct il-

lumination and disturbance regimes in restinga (shore-

line) vegetation at Praia Virgem, municipality of Rio

das Ostras, Rio de Janeiro, Brazil.

MATERIALS AND METHODS

The studied plant material was collected in an area

of restinga vegetation at Praia Virgem, municipality

of Rio das Ostras, Rio de Janeiro, Brazil (22◦31′S ×

41◦56′W). This coastal restinga area occupies about

20 ha on a narrow strip of sand, circa 1 km long, along

the coast (H. Braga, unpublished data). Two distinct veg-

etation formations were identified there (Menezes et al.

2005): Open Palmae Shrub (OPS) and Sandy Strip

Closed Shrub (SSCS), corresponding to the edge and to

the interior of the restinga forest, respectively (Figs. 1a,

b). The region receives 1,100 to 1,300 mm of rainfall

annually, and the climate is mesothermic, sub-humid

to humid, with only a small water deficit during the

year (FIDERJ 1978). The two vegetation formations are

separated by circa 200 m and differ mainly concerning

the amount of sunlight that they receive, their exposure

to the on-shore winds, and in terms of anthropogenic

disturbances, the OPS suffering from frequent pruning.

Samples from three individuals of Mollinedia

glabra were obtained in the vegetation formations (Ta-

ble I). As this species is normally branched near the

base, samples of the SSCS individuals were collected

from the most developed branches at breast height

(1.30 m above ground level), and from the middle of

straight branches of individuals at the OPS site (Ta-

ble I). The wood samples, as suggested by Coradin and

Muñiz (1991), were obtained from the sapwood region,

as the heartwood was rather small. Reproductive ma-

terial was housed at the Herbarium RBR of the Botany

Department of the UFRRJ under the number 401054.

TABLE IList of individuals sampled in both sites, collection sites,

dendrometrics data and record from the wood collection ofInstituto de Pesquisas Jardim Botânico Rio de Janeiro (OPS –

Open Palmae Shrub; SSCS – Sandy Strip Closed Shrub Growing;H = plant height (m), CSH = circumference at sampling height(cm); RBw = record of the wood; SLI = record of the slides).

Indscollection H CSH

RBw SLIsites (m) (cm)

1 OPS 0.6 7.6 8563 2250

2 OPS 1.0 7.5 8564 2251

3 OPS 1.8 7.8 8424 2146

4 SSCS 2.5 8.1* 8425 2147

5 SSCS 3.0 7.8* 8566 2253

6 SSCS 3.5 8.0* 8565 2252

(*) Samples collected at 1.30 m above ground level.

The wood samples from each of the studied indi-

viduals were sectioned along their transversal and lon-

gitudinal planes, at an average thickness of 15μm. The

histological sections were cleared, stained, dehydrated

and mounted as permanent slides. The stain that was

used was a mixture of 1% astra blue and safranin (Bur-

guer and Richter 1991). The permanent slides were

housed at the Xylothek of the Instituto de Pesquisas

Jardim Botânico of Rio de Janeiro (JBRJ) under the


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WOOD ANATOMY OF Mollinedia glabra IN RESTINGA AT RIO DAS OSTRAS, RJ 917

Fig. 1 – a. Aspect of the Open Palmae Shrub (OPS), individuals of Mollinedia glabra (arrows). b. Aspect of the Shrubby

Vegetation (SSCS).

registration numbers indicated in Table I. The wood

fragments were boiled in 50% nitric acid to dissociate

the cellular elements (modified from Occhioni 1948).

The descriptions, counts, and measurements of the cel-

lular elements followed the norms of the IAWA Com-

mittee (1989) and Coradin and Muñiz (1991). The num-

bers of measurements per analyzed individual were: 10,

for frequency and diameter of the vessels elements; 20,

for the ray frequencies; and 25, for other parameters

(Table II).

Quantitative data were obtained by measurements

and counts of 12 parameters observed in the vessels,

fibers, and radial parenchyma in each of the specimens

from the two analyzed vegetation formations. The data

were first tested for normality using the Lilliefors test

(Lilliefors 1967) at an alfa significance level of 0.05.

The data for all of the analyzed parameters were deter-

mined to have normal distributions. The Student t test

was then used to compare the two means of the indepen-

dent data, at an alfa significance level of 0.05, by using

the XLStat version 7.5 software package (Table III).

In order to compare environmental effects on ana-

tomical characters associated with water transport, vul-

nerability, and mesomorphy indices were applied accord-

ing to Carlquist (1975).

Photomicrographs were taken using an Olympus

PM-C35B camera coupled to an Olympus CH30 micro-

scope.


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TABLE IIMean and Standard Deviation of wood anatomical features of Mollinedia glabra

in both plant formations (OPS = Open Palmae Shrub, SSCS = Sandy Strip Closed Shrub).

Mollinedia glabraOPS Standard

SSCS StandardDeviation Deviation

Growth rings distinct indistinct

Porous difuse difuse

Vessels frequency (mm2) 78.5 ± 20.4 57.7 ± 18.7

Vessels lenght (μm) 818.5 ± 76.4 843.3 ± 52.7

Vessels diameter (μm) 38.6 ± 8.0 36.3 ± 5.1

Vessels wall thickness (μm) 3.7 ± 0.2 3.7 ± 0.6

Number of bars

of scalariformPerforation plates 5-15 ± 0.2 7-14 ± 1.1

VesselsRadial multiples 2-5 2-4

Clusters 2-6 2-8

Intervessel Scalariform (μm) 15.2 ± 2.4 13.9 ± 2.0

pitting Opposite (μm) 6.5 ± 1.1 6.4 ± 0.5

Vessel-ray Scalariform (μm) 15.4 ± 7.1 15.7 ± 2.7

pitting Opposite (μm) 8.6 ± 3.5 7.2 ± 0.7

Fibers-tracheids lenght (μm) 1544.1 ± 195.2 1118.3 ± 705.7

Fibers-tracheids diameter (μm) 28.6 ± 2.9 24.9 ± 3.2

Fibers-tracheids lumina (μm) 14.3 ± 1.6 11.8 ± 1.2

Fibers-tracheids wall thickeness (μm) 7.2 ± 0.7 6.5 ± 1.3

Fibers-tracheids pits (μm) 4.3 ± 1.2 3.8 ± 0.85

Gelatinous fibers present present

Rays frequency (linear mm) 3.0 ± 0.6 3.4 ± 0.1

Rays lenght (μm) 677.4 ± 112.4 746.5 ± 156.3

Rays width (μm) 80.6 ± 10.8 80.9 ± 3.3

Rays width (number of cells) 2-8 ± 0.2 2-6 ± 0.5

Sheath cells present present

Prismatic crystals present present

Indice of Vulnerability 0.5 0.6

Indice of Mesomorphy 401.0 522.7

RESULTS AND DISCUSSION

In general, the examined qualitative characters were con-

stant among all studied individuals of M. glabra from

the two restinga formations (Table II), and were consis-

tent with previously published data for the family Mo-

nimiaceae, subfamily Mollinedioideae, and the genus

Mollinedia (Thorne 1974 apud Metcalfe 1987, Barros

et al. 1997, Callado et al. 1997). This species demon-

strates: diffuse porosity; vessel elements that are soli-

tary, in radial groups, or clusters elements, being circular

to angular in cross-section, with short appendices, perfo-

ration plates with few bars (5-15), scalariform to oppo-

site intervessel pitting and vessel-rays pitting predomi-

nantly scalariform; fiber-tracheids septate, ranging from

thin to thick; axial parenchyma extremely rare; multiseri-

ate rays wide (uniseriate rare), heterocellular, composed

of procumbent cells in the central region and marginal

rows of up right and square cells, with the presence of

aggregated and fused rays, sheath cells and prismatic

crystals (Figs. 2-4).

Species of Mollinedia previously studied at Rio

de Janeiro State in an area of Dense Montane Ombrophi-

lous Forest (Barros et al. 1997) demonstrated indistinct


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TABLE IIIStudent t test results for parameters analyzed in both

plant formations (α = 0,05).

Variables t (observed) p

VESSELS

Frequency mm2 4.39 <0.0001*

Lenght 0.37 0.7126

Diameter 1.63 0.1051

Wall thickeness –0.12 0.8997

FIBERS-TRACHEIDS

Lenght 0.46 0.6452

Diameter 4.29 <0.0001*

Lumina 3.87 0.0001*

Wall thickeness 1.65 0.1008

RADIAL PARENCHYMA

Lenght –2.52 0.0124*

Width –0.05 0.9577

Number of cells –0.92 0.3570

Frequency linear mm –2.26 0.0250*

Values marked with (*) are statistically significant (P<0.05).

growth rings. In the studied specimens of M. glabra, the

degrees of distinction of these rings ranged between the

two different formations (Table II), with distinct layers

marked by the thick-walled and radially flattened late-

wood fibers among individuals from the OPS (Fig. 2a).

The formation of growth rings in tropical species

is usually associated with seasonal factors such as

droughts, flooding, changes in the photoperiod, phenol-

ogy, endogenous factors, or in response to stochastic

factors such as insect attacks or diseases, the loss of

branches, or other injuries (Worbes 1989, 1999, Alves

and Angyalossy-Alfonso 2000, Callado et al. 2001,

2004). In the OPS area, the individuals of M. glabra

experience frequent cutting and form latewood, laying

down growth rings that probably represent responses

to these injuries when they cease their radial growth

due to the energetic investments required to form new

branches. Another important characteristic of the in-

dividuals growing in the OPS is the presence of tyloses

with sclerified walls (Fig. 3b). This characteristic can

likewise be considered a response to anthropogenic

disturbances in this formation. Tyloses are observed

in some genera of the family Monimiaceae (Metcalfe

1987), although sclerification of these cells has not been

mentioned previously in the literature. Carlquist (2001)

considered the capacity to form sclerified tilos a valid

taxonomic characteristic.

Among the examined quantitative characteristics,

the two restinga formations only demonstrated signifi-

cant differences in terms of the frequency of the ves-

sel elements, the diameter of the lumen of the fiber-

tracheids, and the length and the frequency of the rays

(Table III). Numerous authors have noted that species

growing in dry environments usually demonstrate in-

creased frequencies of vessels with small diameters, as-

sociating conductive efficiency with conductive safety

(Carlquist 1977, Baas et al. 1983). The individuals of

M. glabra examined in the OPS area demonstrated a

greater frequency of vessel elements than the one re-

ported in SSCS individuals (Table II). In spite of this

variation in their frequency, no significant statistical dif-

ferences were observed in terms of the diameters of

these elements (Tables II and III). Similar results for

the variation of the frequency and diameter of vessel

elements were found for the wood of Xilopia aroma-

tica (Lam.) Mart. growing in cerrado (savanna) areas,

and in plantations of Pinus elliottii Engelm (Luchi et

al. 2005).

The higher frequency of vessel elements associ-

ated with fiber-tracheids that have increased diameters

and lumens can be considered a structural adaptation re-

lated to the water requirements of plants growing in the

OPS site, due to the greater exposure of these plants to

direct sunlight. The fiber-tracheids in M. glabra are sep-

tate (Fig. 3c, d), which is a characteristic typical of the

genus Mollinedia (Metcalfe 1987). Esau (2000) men-

tioned that, in taxa where axial parenchyma is extremely

rare, as in the studied species, the septate fibers are re-

sponsible for the transport and storage of photosynthetic

products.

Although statistical tests did not demonstrate sig-

nificant differences in the lengths of the vessel elements

between the two restinga formations, they did appear

smaller in individuals from the OPS. This characteristic,

together with the smaller numbers of bars on the perfo-

ration plates of the vessel elements (Fig. 2c, Table II),

represents adaptations to drier environments (Wheeler

and Baas 1991, Dickison 2000, Denardi and Marchiori

2005). Reductions in the numbers of bars per perfora-

tion plate in plants growing in xeric environments were


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Fig. 2 – Cross and longitudinal sections of the branches of Mollinedia glabra. a. Growth rings distinct (square),

difuse-porous, vessels clusters (circle) and solitary of OPS (CS). b. Growth rings indistinct or absent, difuse-

porous, vessels paired (circle) and solitary (arrow) of SSCS (CS). c. Scalariform plates (arrow) of OPS (RLS).

d. Intervessel opposite (arrow) of OPS (TLS). Bar = 200μm (a); 100μm (b, c); 50μm (d).

noted for M. glabra (5-15 bars) when compared to other

species of Mollinedia growing in Dense Montane Om-

brophilous Forests (and having from 20 to more than 40

bars) (Barros et al. 1997, Callado et al. 1997).

In spite of the placement of individuals of the

two formations within the mesomorphic range (indices

greater than 200), the results obtained in the present

study corroborate the indices reported by Carlquist

(1977) and indicate the presence of characteristics that

are less mesomorphic and more vulnerable to xylem

hydraulic conductivity in OPS individuals (Table II).

Previous analyses undertaken with the species Molline-

dia gilgiana Perkins, M. marliae A. Peixoto et V. Pe-

reira, and M. salicifolia Perkins in Serra de Macaé de

Cima (Callado et al. 1997) all demonstrated greater

mesomorphic indices (5,969, 1,360 and 6,740, respec-

tively) than those observed here with the specimens of

M. glabra (401 and 522).


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Fig. 3 – Mollinedia glabra wood. a. Details of intervessel pits scalariform to opposite (arrow) of OPS (TLS).

b. Tyloses lightly sclerosed (arrows) and common tyloses (square) of OPS (RLS). c. Fiber-tracheids septate (arrows)

of OPS (TLS). d. Fiber-tracheids (arrows) and presence of prismatic crystal (square) SSCS (CS). Bar = 50μm (a, d);

100μm (b, c).

The increase in the frequency and length of the rays

among the species studied here may be associated with

the increasing diameters of the branches among individ-

uals in the SSCS site (Fig. 4). The greater height of the

rays has been interpreted as being important for the effi-

ciency of radial transport between the pith and the bark

(Catesson 1990, Yáñez-Espinoza et al. 2001).

The results obtained in the present study demon-

strated phenotypic variations in the wood of the spe-

cies being studied, and pointed towards adaptations to

micro-environmental variations during the development

of plants growing in the restinga at Praia Virgem. The

dioecious character of this species and the proximity of

the individual plants favor the exchange of genetic ma-

terial and argue for the existence of a single population

of M. glabra in both vegetation formations.

ACKNOWLEDGMENTS

The authors would like to thank MSc. Henrique do

Nascimento Braga for his help in locating the species

and making the collections; MSc. Alexandre Gabriel

Christo for performing the statistical analyses; the La-

boratório de Botânica Estrutural of the Instituto de Pes-

quisas Jardim Botânico for logistical support; and the

Fundação Carlos Chagas Filho de Amparo à Pesquisa

no Estado do Rio de Janeiro (FAPERJ) for the grant

awarded to the second author. To the anonymous re-

viewers, for their valuable comments.


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Fig. 4 – Longitudinal sections of branches of Mollinedia glabra. a. Upright cells, square and procumbent of

OPS (RLS). b. Upright cells (arrow), square and procumbent of SSCS (RLS). c. Multisseriate rays (?), fusioned

(square), sheath cells (arrow) of OPS (TLS). d. Multisseriate rays (?), sheath cells (black arrow); fiber-tracheids

septate (white arrows) of SSCS (TLS). Bar = 100μm (a; c; d), 400μm (b).

RESUMO

Este trabalho objetiva caracterizar a estrutura anatômica do

lenho de Mollinedia glabra, ocorrente em duas formações

vegetais contíguas na restinga da Praia Virgem, município de

Rio das Ostras, RJ. Essas formações Arbustiva Aberta de Pal-

mae (AAP) e Arbustiva Fechada do Cordão Arenoso

(AFCA) estão sobre cordão arenoso e recebem precipitações

anuais de 1.100-1.300 mm. Foram obtidas amostras do lenho

das duas formações. São características anatômicas gerais da

espécie: elementos de vasos solitários, em arranjos radiais ou

cachos, de seção de circular a angular; placa de perfuração es-

calariforme, com 5-15 barras; parênquima axial escasso; fibro-

traqueídes septadas e raios multisseriados largos, com cristais

prismáticos. A análise estatística evidenciou aumento signi-

ficativo na frequência dos elementos de vasos e aumento no

diâmetro das fibrotraqueídes nos indivíduos de AAP. Essas

características são consideradas adaptações estruturais ao au-

mento da necessidade de água causada por maior exposição

à luz solar. A realização de podas frequentes pode ter relação

com a formação de tilos nesses indivíduos. O aumento signi-

ficativo no comprimento e frequência de raios podem repre-

sentar uma resposta ao maior diâmetro dos indivíduos em

AFCA. Os resultados obtidos sugerem que M. glabra desen-

volve adaptações estruturais às variações micro-ambientais da

restinga durante seu desenvolvimento.

Palavras-chave: Mollinedia glabra, Monimiaceae, restinga,

anatomia do lenho.


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Wood anatomy of Mollinedia glabra (Spreng.) Perkins

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